A peer-reviewed open-access journal ZooKeys 317: An27–52 illustrated (2013) key to the genera of ( Thysanoptera, ) from Iran 27 doi: 10.3897/zookeys.317.5447 Research article www.zookeys.org Launched to accelerate biodiversity research

An illustrated key to the genera of Thripinae (Thysanoptera, Thripidae) from Iran

Majid Mirab-balou1,2, Kambiz Minaei3, Xue-Xin Chen1

1 Institute of Sciences, Zhejiang University, 866 Yuhangtang Road, Hangzhou 310058, China 2 De- partment of Plant Protection, College of Agriculture, Ilam University, Ilam, Iran 3 Department of Plant Pro- tection, College of Agriculture, Shiraz University, Fars, Iran

Corresponding author: Xue-Xin Chen ([email protected])

Academic editor: Laurence Mound | Received 1 May 2013 | Accepted 16 July 2013 | Published 18 July 2013

Citation: Mirab-balou M, Minaei K, Chen X-X (2013) An illustrated key to the genera of Thripinae (Thysanoptera, Thripidae) from Iran. ZooKeys 317: 27–52. doi: 10.3897/zookeys.317.5447

Abstract An illustrated key is provided for the identification of 35 genera of Thripinae (Thysanoptera: Thripidae) from Iran with comments for each . maximi Ananthakrishnan and cerealium Haliday are recorded from Iran for the first time. A checklist is provided of Thripinae recorded from this country.

Keywords Thysanoptera, Thripinae, identification, Iran

Introduction

The Thripidae (Thysanoptera: Terebrantia) at present comprises more than 2000 described , which are classified into four subfamilies, Thripinae, Dendro- thripinae, Sericothripinae, and Panchaetothripinae (Bhatti 1989). The present study follows the interpretation of Thripinae in this classification. Another recent classifica- tion of Terebrantia (Bhatti 2006) recognizes three superfamilies and 12 families for taxa included in the four subfamilies of Thripidae, and the taxa of the earlier subfam- ily Thripinae are included in three families (Chirothripidae, Projectothripidae, and Thripidae) (Bhatti 2006). Thripinae (sensu Bhatti 1989) comprising 1600 species in 230 genera worldwide is the largest subfamily. Members of Thripinae exhibit a wide range of biologies. Many

Copyright Majid Mirab-balou et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 28 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013) species live in flowers, on leaves, some species live in both habitats, particularly the pest species, and a few species are predators. In this subfamily, several genus-groups, which are probably monophyletic, have been recognized, including the genus- group, the genus-group, the genus-group, the genus-group, the Trichromothrips genus-group, the genus-group and the genus-group (Mound and Palmer 1981a, Mound 2002, Masumoto and Oka- jima 2005, 2006, 2007, Mound and Masumoto 2009). Countries of the eastern Mediterranean comprised the most important centre for the early development of human civilization, including the development of the agricultural systems on which so much of mankind depends. In contrast, our knowledge of the natural biological systems of this area has been less actively developed. Despite excellent floristic studies, such as Flora Iranica that now provides an identification system to more than 10,000 plant species, comprehensive studies on the insect fauna of this area are sadly lacking. Iran, in particular, is a bridge between the faunas of the European and Oriental Realms, and this produces considerable difficulties in studying any single group. Keys are available for species of some Iranian genera, such as for the genera of Thrips and Frankliniella genus-groups (Minaei et al. 2007; Mirab-balou and Chen 2011a), and the Megalurothrips genus-group (Mirab-balou and Chen 2011b). Within the 35 genera of Thripinae that are now listed from Iran, many species are widely distributed and their habitats are known. The species of several genera are grass- living, including Agalmothrips, Anaphothrips, , Bregmatothrips, Chirothrips, Collembolothrips, Exothrips, Limothrips, Sitothrips, Sphaeropothrips, Stenchaetothrips, and Stenothrips. On the other hand, many species especially in Thrips and Frankliniella live in various flowers, and these include economic pests of agricultural crops, fruit trees, ornamental plants, greenhouses (Mirab-balou and Chen 2011a, Mirab-balou et al. 2012a). A few are predators, such as species of Scolothrips and Parascolothrips, playing an important role in checking the multiplication of tetranychid mites (Mound 2011b). An annotated bibliography of publications on Thysanoptera of Iran was provided by Bhatti et al. (2009a). However there is not any available key to distinguish the genera. The objective of this paper is to provide an identification key to the 35 genera of Thripinae that can be recognized currently in Iran. Comments are provided for each genus. A checklist of Thripinae known from Iran are also represented here.

Material and methods

For new records, thrips have been prepared and mounted on slides using the method of Mirab-balou and Chen (2010a) and specimens are deposited in the Institute of Insect Sciences, Zhejiang University, Hangzhou, China (ZJUH). All descriptions, measurements and photos were made with a Leica DM IRB microscope, a Leica MZ APO microscope with a Leica Image 1000 system. All measurements are given in micrometers (μm). An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 29

Key to genera of Thripinae from Iran 1 Pronotum without any posteroangular setae longer than discal setae (Fig. 5)....2 – Pronotum with at least one pair of posteroangular or posteromarginal setae longer than discal setae (Figs 1–4, 6)...... 5 2 Antennal segments III and IV each with a simple sensorium...... 3 – Antennal segments III and IV each with a forked sensorium...... 4 3 Antennae 9-segmented (cf. Fig. 26); apterous or macropterous; abdominal sternites without discal setae; tarsi 2-segmented; male with a transverse pore plate on abdominal sternites III–VII...... Agalmothrips – Antennae 6- or 8-segmented (Fig. 29); apterous; abdominal sternites with or without discal setae; tarsi 1- or 2-segmented (tarsi 2-segmented if antennae 8-segmented, in A. stylifer Trybom); male without pore plate on abdominal sternites...... Aptinothrips 4 Median pair of setae (S1) on abdominal tergites II–VIII shorter than distance between their bases (Fig. 12); abdominal tergite VIII with comb at posterior margin...... Anaphothrips – Median pair of setae (S1) on abdominal tergites II–VIII longer than distance between their bases; abdominal tergite VIII without comb at posterior mar- gin...... Rubiothrips 5 Abdominal tergites V–VIII each with a pair of lateral ctenidia...... 6 – Abdominal tergites V–VIII without ctenidia...... 13 6 Abdominal tergite VIII with ctenidium situated antero-lateral to spiracle....7 – Abdominal tergite VIII with ctenidium situated posteromesad of spiracle.... 9 7 Pronotum anterior margin without long setae (cf. Fig. 1); mesosternum with- out spinula; maxillary palp 2-segmented (cf. Fig. 20)...... Sitothrips – Pronotum anterior margin with 1 or 2 pairs of setae much longer than discal setae (Fig. 6); mesosternum with spinula (cf. Fig. 15); maxillary palp 3-seg- mented (cf. Fig. 22)...... 8 8 Fore tarsus with a small tooth at apex (Fig. 23); ocellar setae pair III inserted between posterior ocelli; abdominal sternite VII of female with S1 setae in- serted ahead of posterior margin (cf. Fig. 36)...... Kakothrips – Fore tarsus without tooth (Fig. 21); ocellar setae pair III arising at a level ahead of posterior ocelli (Fig. 9); abdominal sternite VII of female with S1 setae arising at posterior margin (Fig. 38)...... Frankliniella 9 Mesothoracic sternopleural sutures absent; antennal segment II without seta basad of campaniform sensillum...... Sphaeropothrips – Mesothoracic sternopleural sutures present; antennal segment II with dorsal seta basad of campaniform sensillum...... 10 10 Prosternal basantra with several small setae; abdominal tergites at posterior margin with large triangular teeth (Fig. 45)...... Microcephalothrips – Prosternal basantra without setae; posterior margin of abdominal tergites without teeth; head not distinctly smaller than pronotum...... 11 30 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

11 Postocular setae pair II inserted far back of the others; abdominal tergites III–V with three setae arranged straight line along the lateral margin...... Stenothrips – Postocular setae pair II not displaced to the posterior although it is often slightly behind setal row; abdominal tergites III–V with three setae arranged not in straight line, median setae far from the lateral margin...... 12 12 Ocellar setae pair II longer than ocellar setae pair III...... Stenchaetothrips – Ocellar setae pair II not longer than ocellar setae pair III (Fig. 10)...... Thrips 13 Spinula absent on mesosternum...... 14 – Spinula present on mesosternum (Fig. 15)...... 18 14 Abdominal tergite X of female with a pair of prominent thorn-like setae (Fig. 8); pronotum with a pair of well developed posteroangular setae. [Male apter- ous, with a pair of short stout setae medially on tubercles on abdominal ter- gite IX (Fig. 44)]...... Limothrips – Abdominal tergite X of female without such thorn-like setae (Fig. 42); prono- tal posteroangular setae variable...... 15 15 Abdominal sternite VII of female with posteromarginal setae S1 and S2 aris- ing closer to each other than to setal pair S3...... Exothrips – Abdominal sternite VII of female with posteromarginal setae arising equidis- tant from each other...... 16 16 Abdominal tergites with median pair of campaniform sensilla close to the posterior margin (Fig. 7); abdominal sternite II with three pairs of posteromarginal setae. [Male apterous, and bicolored yellow and brown (Fig. 35)]...... Bregmatothrips – Abdominal tergites with median pair of campaniform sensilla arising at an- terior third (cf. Figs 14, 46); abdominal sternite II with two pairs of postero- marginal setae...... 17 17 Pronotum broadly rectangular, with 3 pairs of posteromarginal setae; anten- nal segment II not prolonged laterally; fore tarsus with a curved tooth (Fig. 18); maxillary palp 2-segmented...... Collembolothrips – Pronotum trapezoidal, with 5–8 pairs of posteromarginal setae (Fig. 1); an- tennal segment II usually projecting laterally (Fig. 30); fore tarsus without tooth (Fig. 16); maxillary palp 3-segmented...... Chirothrips 18 Metasternum with prominent spinula...... 19 – Metasternum without spinula...... 24 19 Antennae 6-segmented (cf. Fig. 29)...... Drepanothrips – Antennae 7- or 8-segmented (Figs 27–28)...... 20 20 Antennae 7-segmented, VII slightly longer than VI; tarsi 1-segmented (cf. Fig. 17)...... Parascolothrips – Antennae 8-segmented; tarsi 2-segmented...... 21 21 Pronotum with six pairs of very long setae (Fig. 2)...... Scolothrips – Pronotum with no more than two pairs of elongate setae...... 22 22 Abdominal tergites without numerous microtrichia occupying lateral thirds, rarely with a few microtrichia near lateral margins; maxillary palp 2-segment- ed (cf. Fig. 20)...... Psilothrips An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 31

– Abdominal tergites with lateral thirds fully covered with numerous microtri- chia (Fig. 37); maxillary palp 3-segmented (cf. Fig. 22)...... 23 23 Pronotum with two pairs of prominent posteroangular setae; antennal seg- ment I with pair of dorsoapical setae; males of some species with antennal segment VI three times as long as segment V (Fig. 40)...... Mycterothrips – Pronotum with closely spaced lines of sculpture, without long setae, or only one pair of prominent posteroangular setae (Fig. 4); antennal segment I with- out dorsoapical setae; antennae not sexually dimorphic...... Scirtothrips 24 Spiracles on abdominal tergite VIII with area of specialised sculpture extending to antecostal ridge...... Chaetanaphothrips – Spiracles on tergite VIII without extensive area of specialised sculpture.....25 25 Head with 4 or more pairs of ocellar setae...... Florithrips – Head with 2 or 3 pairs of ocellar setae ...... 26 26 Setae on fore wing first and second veins very long, longest seta twice as long as wing width...... Euphysothrips – Setae on fore wing first and second veins shorter, longest seta scarcely as long as wing width...... 27 27 Pronotum with four pairs of setae well-developed; antennae 9-segmented...... Ficothrips – Pronotum with one or two pair of posteroangular setae well-developed, with no long anteromarginals and anteroangulars setae...... 28 28 Pronotum with two pairs of posteroangular setae well-developed...... 29 – Pronotum with one pair of posteroangular setae...... 33 29 Head with only one pair of anteocellar setae (ocellar seta pair I absent) (cf. Fig. 10)...... Taeniothrips – Head with two pairs of anteocellar setae (ocellar seta pair I present) (cf. Figs 9, 11)...... 30 30 Sensorium on antennal segment VI with elongate base (Fig. 31); fore tibia usually with 1 or 2 claw-like processes at apex; fore tarsus often with 1 or 2 small tubercles (Fig. 19)...... – Base of sensorium on antennal segment VI not elongate; fore tibia without tubercles...... 31 31 Fore wing first vein with nearly complete row of setae and short interval before two distal setae (Fig. 34); clavus with 4 or 5 veinal setae...... Megalurothrips – Fore wing upper vein with setal row broadly interrupted, with 1+2 distal setae (cf. Fig. 33); clavus usually with 5 veinal setae...... 32 32 Antennal segment I without pair of dorsoapical setae; male with a single oval or circular pore plate on each of abdominal sternites III–VII (Fig. 43)...Tenothrips – Antennal segment I with a pair of dorsal apical setae (Fig. 32); male with nu- merous small pore plates on abdominal sternites III–VII (Fig. 41)...Pezothrips 33 Antennae 9-segmented...... Eremiothrips [in part] – Antennae 7- or 8-segmented...... 34 32 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

34 Abdominal tergites III–VI with lines of sculpture medially; sternites without discal setae...... – Abdominal tergites without lines of sculpture medially; sternites with or without discal setae (Fig. 39)...... 35 35 Head longitidunaly striate behind eyes; abdominal sternite II with 0–4 discal setae...... Tamaricothrips – Head weakly striate behind eyes; abdominal sternite II without discal setae...... Eremiothrips [in part]

Agalmothrips Priesner

Remarks. Included here only from descriptions, the sole species in this genus, A. par- viceps Priesner, was described from Sudan (Priesner 1964), subsequently reported and re-characterized from India (Bhatti 1978). It was reported from Iran by zur Strassen (2003b) based on two males and eight females collected on Kochia sp. (Chenopodiace- ae), in Ahwaz region (Khuzestan province). Agalmothrips was included in the Anapho- thrips genus-group (Mound and Masumoto 2009).

Anaphothrips Uzel

Remarks. This genus includes 79 species in the world (ThripsWiki 2013), and many of these are grass-living. In Iran, only A. obscurus (Müller) and A. sudanensis Trybom have been reported (Bhatti et al. 2009a). Recently, the male of the widely distributed species A. obscurus is described only from Iran (Mirab-balou and Chen 2010b).

Aptinothrips Haliday

Remarks. The four species included in this genus (Mirab-balou et al. 2011a) are all apterous. Three of these are known from Iran: A. elegans Priesner, A. rufus (Haliday) and A. stylifer Trybom (Bhatti et al. 2009a). The genus is included in the Anaphothrips genus-group (Mound and Masumoto 2009).

Bregmatothrips Hood

Remarks. This is a common genus of grass-living species in tropical and subtropical ar- eas (Mound and Marullo 1996; Mound 2011a), with nine species worldwide (Mound An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 33

Figures 1–8. Thripinae genera.1 Chirothrips aculeatus, pronotum 2 Scolothrips longicornis head and pro- notum 3–6 pronotum 3 Limothrips angulicornis 4 Tenothrips frici 5 Aptinothrips stylifer 6 Frankliniella oc- cidentalis, 7 Bregmatothrips bournieri abdominal tergite VI 8 Limothrips angulicornis, abdominal tergite X. 34 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

2011a, ThripsWiki 2013) of these B. bournieri Pelikán is found in Iran (Pelikán 1988; Bhatti et al. 2009a). This genus is closely related to Sorghothrips by having antennal segment I with paired median dorsoapical setae, and the abdominal tergites with po- steromarginal craspeda and the median campaniform sensilla close to the posterior margin (Masumoto and Okajima 2006).

Chaetanaphothrips Priesner

Remarks. This is an Asian leaf-living genus (Pitkin 1977), and now includes 20 species (ThripsWiki 2013). Several of these are widespread around the world, and are con- sidered pests on their host plants. C. theiperdus is a pest of tea in Java and Peninsular Malaysia; C. leeuweni (Karny), C. orchidii (Moulton) and C. signipennis (Bagnall) are pests of Bananas in some tropical countries (Kudô 1985, Mound and Marullo 1996); C. orchidii is a most widely distributed polyphagous species and is also recorded as a pest of some ornamental plants in greenhouses in Europe and North America (Kudô 1985, Mound and Ng 2009). Species of this genus may be distinguished from other Thripinae by the presence of modified spiracles on abdominal tergite VIII. This genus was reported from Iran by Esmaili (1983), based on an undetermined species (Chaet- anaphothrips sp.) noted as pest of citrus fruits in Iran.

Chirothrips Haliday

Remarks. Species of this genus breed only in the flowers of grasses and are readily recognized from the shape of the head, pronotum, fore legs and antennae. This genus includes 53 species in the world (ThripsWiki 2013). Six species in Chirothrips and two species in Agrostothrips Hood have been reported from Iran (Bhatti et al. 2009a) but subsquentley Minaei and Mound (2010) considered Agrostothrips as a synonym of Chirothrips and five species and one species-group (manicatus group) have been recognized in this country. A key also to the Chirothrips species from Iran was given by Minaei and Mound (2010).

Chirothrips maximi Ananthakrishnan

Remarks. This species is here recorded from Iran for the first time, based on one apterous male. Hamedan province: Hamedan, Qahavand, from Salvia nemorosa (La- miaceae), 20.viii.2010, coll. M. Mirab-balou (in ZJUH). The male of this species is easily distinguished from other species of Chirothrips by having small pore plates on abdominal sternite III and IV. An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 35

Figures 9–15. Thripinae genera. 9–11 Head, 9 Frankliniella occidentalis 10 Thrips alliorum 11 Teno- thrips frici, 12–14 Abdominal tergite VII 12 Anaphothrips obscurus 13 Rubiothrips vitis 14 Chirothrips ac- uleatus, abdominal tergites VII–VIII, 15 Megalurothrips distalis, Meso- and metasterna, showing spinula. 36 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

Collembolothrips Priesner

Remarks. This genus comprises two grass-living species (ThripsWiki 2013) of which C. mediterraneus Priesner has been reported from Iran (Bhatti et al. 2009a). It is easi- ly distinguished from other thripine in Iran by the absence of a spinula on both the meso- and metathoracic furca, also the lack of ocelli and wings, and the 2-segmented maxillary palps.

Drepanothrips Uzel

Remarks. This monobasic genus is included in Scirtothrips genus-group (Mound and Palmer 1981a; Masumoto and Okajima 2007). D. reuteri Uzel is easily recognized from other Iranian genera by precense of the 6-segmented antennae. It is known as a pest of grapes in some areas (Mound et al. 1976; Masumoto 2010).

Eremiothrips Priesner

Remarks. This genus with 18 species (ThripsWiki 2013) is included in Anaphothrips genus-group (Mound and Masumoto 2009). Bhatti et al. (2003) published a catalogue of the Eremiothrips in Iran with key for separating the nine species of this genus. Rec- ognition of species of Eremiothrips based on females is extremely difficult, since the general appearance and fine structural details of females are very similar in related spe- cies. However, males can be assigned readily to species because of distinctive structural features (Bhatti et al. 2003). In Iran, the genus includes 12 species (Bhatti et al. 2009b, Ramezani et al. 2009, Minaei 2012a) and a key to Iranian Eremiothrips based on the male sex is available (Minaei 2012a).

Euphysothrips Bagnall

Remarks. This genus includes two species (ThripsWiki 2013). Both are known from India but E. minozzii Bagnall has been reported from Iran (Bhatti et al. 2009a). The fore wing chaetotaxy is unique, with very long setae on the veins (Mound and Ng 2009).

Exothrips Priesner

Remarks. This genus comprises 18 grass feeding species (ThripsWiki 2013), and is represented in Iran by E. redox Bhatti (Bhatti et al. 2009a). Bhatti (1975) provided an identification key to 10 species from India, and indicated that these are associated with Poaceae. An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 37

Figures 16–25. Thripinae genera. 16–19 Fore tibia and tarsus, 16 Chirothrips aculeatus 17 Limothrips ce- realium 18 Collembolothrips mediterraneus 19 Odontothrips loti, 20 Limothrips cerealium, maxillary palps 21 Frankliniella occidentalis, fore tarsus 22 Megalurothrips distalis, maxillary palps 23–24 Fore tibia and tarsus 23 Kakothrips pisivorus 24 Sitothrips arabicus 25 Bregmatothrips bournieri, mesonotum and metascutum. 38 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

Ficothrips Minaei

Remarks. This genus originally described from Iran with only one species,F. moundi (Minaei 2012b). This monobasic genus is superficially similar to Scolothrips or Paras- colothrips. Morphologically, Ficothrips is interesting because simultaneously bears two charchter states that have been evolved rarely in Thripidae: nine antennal segments and severall long setae on the pronotum. The species was collected on fig leaves infested by Eotetranychus hirsti Pritchard and Baker (Acari, Tetranychidae) and the low density of species suggest that it may be a predator on that mite (Minaei 2012b).

Florithrips Bhatti

Remarks. This genus includes two species in the world (ThripsWiki 2013), of these F. traegardhi was recorded from Iran on the base of a few specimens collected on corn and wheat in Khozestan Province (Ramezani et al. 2012). Leaf damage to cereal crops is recorded for this species (Mound and Kibby 1998).

Frankliniella Karny

Remarks. This is a large genus of about 230 species, 90% of which are from Neotropics (Mound and Marullo 1996), with only five species recorded from Iran (Mirab-balou and Chen 2011a). The genus can be recognized by having ctenidia anterolateral to each spiracle on abdominal tergite VIII, presence of five pairs of long pronotal setae, and a complete setal row on both upper and lower veins of the fore wings (Mound and Marullo 1996). A key to Iranian species is available in Mirab-balou and Chen (2011a). The species of this genus recorded from Iran are most associated with the family Rosaceae (Mirab-balou and Chen 2011a). The reports of two species from Iran have not been ac- cepted (Bhatti and zur Strassen 2009): F. cephalica (D.L. Crawford) and F. tritici (Fitch).

Kakothrips Williams

Remarks. This genus includes seven species (ThripsWiki 2013), of which three are recorded from Iran (Mirab-balou and Chen 2011a). Kakothrips is included in the Frankliniella genus- group (Mound and Palmer 1981a; Mirab-balou and Chen 2011a). This genus is distin- guished from Frankliniella by moderately developed ctenidia laterally on abdominal tergites VI–VII whereas well-developed in Frankliniella, and the pronotum lacks a pair of minor setae medially on the posterior margin. In addition, males of Kakothrips species have a pair of stout tubercles laterally on tergite VIII, whereas none of the Frankliniella species has such structures (Moritz et al. 2001). Zur Strassen (2003a) provided a key to seven species from Europe; and a key to the three species in Iran is available in Mirab-balou and Chen (2011a). An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 39

Figures 26–34. Thripinae genera. 26–32 Antennae, 26 Anaphothrips obscurus 27 Pezothrips kellyanus 28 Microcephalothrips abdominalis 29 Aptinothrips elegans 30 Arorathrips mexicanus 31 Odontothrips con- fusus, sensoria on segment VI 32 Trichromothrips sp., dorsal apical setae on segment I 33–34 Fore wing 33 Taeniothrips inconsequens 34 Megalurothrips distalis. 40 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

Limothrips Haliday

Remarks. This western Palaearctic genus includes eight species (ThripsWiki 2013), of which four species have been reported from Iran (Bhatti et al. 2009a). Females of this ge- nus can be distinguished easily from other genera of Thripinae by having abdominal ter- gite X with short, stout, spine-like median setae (zur Strassen 2003a, Masumoto 2010). Limothrips cerealium Haliday is here recorded from Iran for the first time, based on one apterous female: Azarbaijan-e-Sharghi province: Tabriz, Miyaneh, from harvested wheat, Triticum aestivum L. (Poaceae), 26.vi.2009, M. Mirab-balou, (in ZJUH). This species is distinguished from other species by tarsi 1-segmented, absence of ocelli, and antennal segments III and IV with simple sensoria. This European pest of cereal crops is now widespread throughout the temperate and subtropical areas of the world. Adults are usuallymacropterous, but apterae have been recorded from Sardinia (Karny 1914) and Corsica (Mound and Palmer 1973), and now from Iran.

Megalurothrips Bagnall

Remarks. Thirteen species are included in this genus (ThripsWiki 2013), all breeding in the flowers of Fabaceae some as pests of cultivated legumes (Masumoto 2010). Recently, M. distalis (Karny) was recorded from Iran (Mirab-balou and Chen 2011b). The members of this genus all have a pair of dorso-apical setae on the first antennal segment, and abdominal tergite VIII with many scattered microtrichia anterior to the spiracles (Mound and Ng 2009).

Microcephalothrips Bagnall

Remarks. This monobasic genus is included in Thrips genus-group (Mound and Palmer 1981a; Mirab-balou and Chen 2011a), from which it can be distinguished by the key above. M. abdominalis lives in the flowers of various Asteraceae, particularly sunflower,Helianthus annuus (Palmer 1992; Mound and Marullo 1996). The macrop- terous morph is common in Iran. Recently one micropterous male was collected on grasses from Ilam Province, and is firstly recorded of micropterous morph for Iran.

Mycterothrips Trybom

Remarks. The 27 known species of Mycterothrips (ThripsWiki 2013) are leaf-feeding thrips, and some of them are associated with agricultural crops (Masumoto and Oka- jima 2006). An identification key to five species from Taiwan was provided by Wang (1999), a key to seven species has been provided by zur Strassen (2003a), and a key to 27 species by Masumoto and Okajima (2006). Up to now, four species have been re- corded from Iran, with two new species described recently (Mirab-balou et al. 2011b). An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 41

Figures 35–40. Thripinae genera. 35 Bregmatothrips bournieri, apterous male 36 Pezothrips kellyanus, abdominal sternite VII 37 , abdominal tergites V–VI 38 Frankliniella tenuicornis; ab- dominal sternite VII 39 Microcephalothrips abdominalis, abdominal sternites V–VI 40 Mycterothrips con- sociatus, antennal segment IV–VIII, male. 42 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

Odontothrips Amyot & Serville

Remarks. The 31 species of this genus (ThripsWiki 2013) are typical flower-living thrips, and most are associated with plants of the family Fabaceae. They can cause slight damage to the flowers, but only O. confusus is an important pest (Pitkin 1972). Megalurothrips peculiaris which sensorium base is similar to Odontothrips, and recorded from India and Bangladesh. This genus is included in the Megalurothrips genus-group (Mound and Palmer 1981a) and four species have been recorded from Iran (Mirab- balou and Chen 2011b). O. confusus Priesner is widely distributed in Iran, and popula- tions are sometimes high on alfalfa, Medicago sativa.

Oxythrips Uzel

Remarks. This genus includes 50 species (ThripsWiki 2013), of which five species are known from Iran (Bhatti et al. 2009a). Oxythrips is included in Anaphothrips genus- group (Mound and Masumoto 2009).

Parascolothrips Mound

Remarks. This monobasic genus with one predatory species, P. priesneri Mound, has been reported from Iraq (Mound 1967, ThripsWiki 2013). Mound (1967) described the species from Iraq with three pairs of posteromarginal setae on sternites II–VI, but in Iranian specimens, there are three pairs of setae on sternite II and four pairs on sternites III–VI.

Pezothrips Karny

Remarks. This genus is placed in theMegalurothrips genus-group (Mound and Palmer 1981a; Mirab-balou and Chen 2011b) and currently includes 10 species (Mirab-balou and Tong 2012). Only one species, P. bactrianus (Pelikan), has been reported from Iran (Mirab-balou and Chen 2011b).

Psilothrips Hood

Remarks. Included here only from descriptions, this genus includes five species in the world (ThripsWiki 2013) of whichP. bimaculatus (Priesner) has been reported from Iran (Bhatti et al. 2009a). An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 43

Figures 41–46. Thripinae genera. 41 Pezothrips kellyanus, pore plates on abdominal sternites V–VII 42 Pezothrips kellyanus, abdominal tergite X 43 Tenothrips frici, pore plate on abdominal sternite V 44 Limothrips angulicornis, short stout setae medially on tubercles on abdominal tergite IX 45 Microcephalo- thrips abdominalis, abdominal tergites VI–VIII 46 Chirothrips molestus, abdominal tergites III–IV. 44 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

Rubiothrips Schliephake

Remarks. This genus includes seven species (ThripsWiki 2013), and one of these has been recorded from Iran (Bhatti et al. 2009a). The genus is included in the Anapho- thrips genus-group (Mound and Masumoto 2009), and is distinguished from Anapho- thrips by having abdominal tergites with median pair of setae (S1) longer than inter- distance (zur Strassen 2003a).

Scirtothrips Shull

Remarks. The genus Scirtothrips currently includes 103 species (ThripsWiki 2013), and two of these have been recorded from Iran (Bhatti et al. 2009a). It includes sev- eral important pest species (Mound and Palmer 1981b, Mirab-balou et al. 2012b). The report of S. citri (Moulton) from Iran has not been accepted (Bhatti and zur Strassen 2009).

Scolothrips Hinds

Remarks. Species of the genus Scolothrips are well known as predators of mites on the leaves of plants. The genus is easy to recognize by the presence of six pairs of very long setae on the pronotum, and the fore wings with dark bands, although recognition of species within the genus has remained difficult (Mound 2011b). Currently 16 species are recognized (ThripsWiki 2013), of which three are reported from Iran (Bhatti et al. 2009a); but the report of S. sexmaculatus (Pergande) from Iran has not been accepted (Bhatti and zur Strassen 2009).

Sitothrips Priesner

Remarks. This genus includes four species (ThripsWiki 2013) of which S. arabicus Priesner has been recorded from Iran (Bhatti et al. 2009a, Mirab-balou and Chen 2011a). This genus is included in the Frankliniella genus-group (Mound and Palmer 1981a). It is distinguished from other members of this group by lack of long setae on the anterior margin of the pronotum, the meso- and metathoracic furcae both without a spinula, the maxillary palps 2-segmented, and the fore tarsi with two small teeth (one at the apex of the tarsus, the other at the end of the first segment) (Mirab-balou and Chen 2011a). A key to three species of Sitothrips is available in zur Strassen (2003a). In some parts of Iran, like as Golestan province, S. arabicus is largely present on wheat and barley (Alavi et al. 2007) An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 45

Sphaeropothrips Priesner

Remarks. This monotypic genus includes a single grass-living species, S. vittipennis (Bagnall) that was recorded from Iran by Minaei et al. (2007). It is included in Thrips genus-group (Mound and Palmer 1981a).

Stenchaetothrips Bagnall

Remarks. This genus includes 35 species (ThripsWiki 2013) and it is associated with the plant family Poaceae. S. biformis is widely distributed, and was recorded from Iran by Mirab-balou and Chen (2011a). The genus is included in Thrips genus-group (Mound and Palmer 1981a; Mirab-balou and Chen 2011a) and is closely related to Thrips, but it can be distinguished by the length of ocellar setae II as in the above key, and the metanotum is generally longitudinally striate (Mound and Ng 2009).

Stenothrips Uzel

Remarks. The only species in this genus, S. graminum Uzel, was recorded from Iran by Minaei et al. (2007). It is a member of Thrips genus-group (Mound and Palmer 1981a).

Taeniothrips Amyot & Serville

Remarks. This genus includes 25 species in the world (Mound et al. 2012; Thrip- sWiki 2013). T. inconsequens (Uzel) is the only species recorded from Iran (Bhatti et al. 2009a).

Tamaricothrips Priesner

Remarks. Only one species is placed in this genus, is also recorded from Iran (Bhatti et al. 2009a). This species is possibly more widespread in association with Tamarix spe- cies (zur Strassen 2003a). The genus is included in Anaphothrips genus-group (Mound and Masumoto 2009).

Tenothrips Bhatti

Remarks. This genus includes 19 flower-living species in the world (ThripsWiki 2013), of which T. frici (Uzel), is widespread around the world in warm temper- ate areas (Mound and Marullo 1996; Mirab-balou and Tong 2013). Bhatti (2003) 46 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013) reviewed this genus, listing 19 species. Four species are reported from Iran (Bhatti et al. 2009a), especially on the plant family Asteraceae, and T. frici is widely distributed in this country.

ThripsLinnaeus

Remarks. This genus includes about 280 species in the world and is the largest genus in the Thripinae (ThripsWiki 2013), and includes 29 species in Iran (Mirab-balou et al. 2012a; Minaei 2012c). The genus shows extensive diversity in most parts of the world except the Neotropical region, and includes many species of economic importance (Bhatti 1980). Sev- eral species of Thripsare considered crop pests in various parts of the world, such as T. angus- ticeps Uzel, T. flavusSchrank, T. hawaiiensis (Morgan), T. meridionalis Priesner, and T. ta- baci Lindeman (Moritz et al. 2001). The latter species is well known as the most important pest of onion crops, greenhouses and ornamental plants in Iran (Mirab-balou and Chen 2011a, Mirab-balou et al. 2012b), and is a vector of some Tospovirus diseases on plants.

Acknowledgements

We are grateful to Prof. J.S. Bhatti of New Delhi, India for his critical comments and suggestions on a draft of this paper and for supplying some useful references; to Dr. Laurence A. Mound of CSIRO Ecosystem Sciences, Canberra, Australia for supplying some references and his critical comments; and to Prof. Xiao-li Tong of South China Agricultural University, Guangzhou, China for supplying some references. The authors are grateful to anynomous referees for their comments and criticisms.

References

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Appendix Checklist of Thripinae known from Iran

Agalmothrips parviceps Priesner Anaphothrips obscurus (Müller) Anaphothrips sudanensis Trybom Aptinothrips elegans Priesner Aptinothrips rufus (Haliday) Aptinothrips stylifer Trybom Bregmatothrips bournieri Pelikan Chaetanaphothrips sp. Chirothrips aculeatus Bagnall Chirothrips africanus Priesner Chirothrips atricorpus Girault Chirothrips kurdistanus zur Strassen Chirothrips manicatus (Haliday) Chirothrips maximi Ananthakrishnan Chirothrips meridionalis Bagnall Chirothrips molestus Priesner Chirothrips pallidicornis Priesner Collembolothrips mediterraneus Priesner Drepanothrips reuteri Uzel Eremiothrips antilope (Priesner) Eremiothrips arya (zur Strassen) Eremiothrips bhattii Minaei Eremiothrips dubius (Priesner) Eremiothrips efflatouni (Priesner) Eremiothrips farsi Bhatti and Telmadarraiy Eremiothrips shirabudinensis (Jaknontov) Eremiothrips similis Bhatti Eremiothrips taghizadehi (zur Strassen) Eremiothrips tamaricis (zur Strassen) Eremiothrips varius (Bhatti) Eremiothrips zurstrasseni Bhatti, Bagheri, and Ramezani Euphysothrips minozzii Bagnall Exothrips redox Bhatti Ficothrips moundi Minaei Florithrips traegardhi (Trybom) Frankliniella intonsa (Trybom) Frankliniella occidentalis (Pergande) Frankliniella pallida (Uzel) (Trybom) An illustrated key to the genera of Thripinae( Thysanoptera, Thripidae) from Iran 51

Frankliniella tenuicornis (Uzel) Kakothrips dentatus Knechtel Kakothrips pisivorus (Westwood) Kakothrips priesneri Pelikan Limothrips angulicornis Jablonowski Limothrips cerealium Haliday Limothrips denticornis (Haliday) Limothrips schmutzi Priesner Limothrips transcaucasicus Savenko Megalurothrips distalis (Karny) Microcephalothrips abdominalis (Crawford) Mycterothrips consociatus (Targioni-Tozzetti) Mycterothrips hamedaniensis Mirab-balou, Shi and Chen Mycterothrips latus (Bagnall) Mycterothrips salicis (Reuter) Mycterothrips tschirkunae (Yakhontov) Mycterothrips weii Mirab-balou, Shi and Chen Odontothrips confusus Priesner Odontothrips meliloti Priesner Odontothrips loti (Haliday) Odontothrips phlomidinus Priesner Oxythrips claripennis Priesner Oxythrips halidayi Bagnall Oxythrips retamae (Priesner) Oxythrips ulmifoliorum (Haliday) Oxythrips wiltshirei Priesner Parascolothrips priesneri Mound Pezothrips bactrianus (Pelikan) Psilothrips bimaculatus (Priesner) Rubiothrips vitis (Priesner) Scirtothrips citri (Moulton) Scirtothrips mangiferae Priesner Scolothrips latipennis Priesner Scolothrips longicornis Priesner Scolothrips rhagebianus Priesner Sitothrips arabicus Priesner Sphaeropothrips vittipennis (Bagnall) Stenchaetothrips biformis (Bagnall) Stenothrips graminum Uzel Taeniothrips inconsequens (Uzel) Tamaricothrips tamaricis (Bagnall) Tenothrips discolor (Karny) Tenothrips frici (Uzel) 52 Majid Mirab-balou et al. / ZooKeys 317: 27–52 (2013)

Tenothrips latoides (Pelikán) Tenothrips reichardti (Priesner) Thrips alaviiMirab-balou, Tong and Chen Thrips alliorum(Priesner) Thrips angusticepsUzel Thrips atratusHaliday Thrips australis(Bagnall) Thrips coloratusSchmutz Thrips dubiusPriesner Thrips euphorbiaeKnechtel Thrips flavusSchrank Thrips fraudulentus(Priesner) Thrips fuscipennisHaliday Thrips hawaiiensis(Morgan) Thrips iranicusYakhontov Thrips majorUzel Thrips mareoticus(Priesner) Thrips meridionalis(Priesner) Thrips minutissimusLinnaeus Thrips nigropilosusUzel Thrips pelikaniSchliephake Thrips physapusLinnaeus Thrips pillichiPriesner Thrips pistaciaeYakhontov (Morison) Thrips tabaciLindeman Thrips treherneiPriesner Thrips trybomi(Karny) Thrips verbasci(Priesner) Thrips vuilleti(Bagnall) Thrips vulgatissimusHaliday